Offspring recognition by mother crab spiders with extreme maternal care

Maternal care is provided by several spider species, but there are no reports of mother spiders recognizing their young, which suggests that maternal care can be exploited by unrelated individuals. Diaea ergandros, a crab spider with extreme, sacrificial maternal care, does accept unrelated spiderlings (ca. 43.9% of spiderlings) into its nest in areas of high nest density. However, a field and a laboratory experiment with mother spiders and natural and adoptive spiderlings demonstrated that mothers did recognize their own offspring. Recognition was not expressed in survival as adopted (unrelated) spiderlings had similar survival rate to that of natural offspring. Instead it was displayed in growth; mother D. ergandros caught large prey items for their own offspring, but not for adopted spiderlings, and so natural offspring grew more than adopted spiderlings. Also, mothers produced trophic oocytes, which are important for the sacrificial care that influences spiderling survival, only when they lived with their own offspring.     

Adaptive advantages of matriphagy in the foliage spider, Chiracanthium japonicum (Araneae: Clubionidae)

Young of the Japanese foliage spider, Chiracanthium japonicum, show matriphagy, whereby they consume their own mothers before dispersal. By removing mothers in the laboratory, I examined the importance of this sacrificial habit for offspring survival and dispersal behavior. Spiderlings that cannibalized their mothers gained weight more than threefold and dispersed from their breeding nests after molting into the third instar. The third-instar spiderlings had relatively longer legs than the previous instars and appeared to be more adapted to a solitary hunting life style. On the other hand, most spiderlings separated from their mothers could not molt into the third instar and dispersed significantly earlier than those with matriphagy. Furthermore, the lack of matriphagy decreased the survival rate of predispersal spiderlings. These results showed that matriphagy of C. japonicum has a great advantage in allowing offspring to disperse at a more developed and active instar. Received: October 23, 2000 / Accepted: March 15, 2001     

Spiderling sex ratio and maternal investment in the bolas spider Mastophora cornigera (Araneae,Araneidae)

Male Mastophora cornigera exit egg sacs as adults, which allowed us to determine spiderling sex ratios and patterns of maternal investment in this species. We collected 15 egg sacs produced by seven mothers, which yielded 1945 emergent spiderlings which were sexed, 1850 of which were weighed. Two emergent broods were significantly male and female biased and were unaffected by pre-emergence mortality. The weights of male and female spiderlings differed in eight broods, with males and females being heavier in four cases each. Five of these broods were derived from multiple egg sac sets produced by one mother, and in each case, the total mean male and female spiderling weights for all broods in a set were biased in the same direction as the biased brood(s) within that set. Mean emergent spiderling weight was independent of brood size and sex ratio for both males and females. Despite such independence, sex allocation in M. cornigera can favor sons, daughters, or both equally, and by numbers, by weight, or both at once. The proximate mechanisms and adaptive significance of such variability is unknown. We also review evidence for gender-biased allocations in arachnid offspring and suggested mechanisms for their applicability to M. cornigera.     

The allometry of CNS size and consequences of miniaturization in orb-weaving and cleptoparasitic spiders

Allometric studies of the gross neuroanatomy of adults from nine species of spiders from six web-weaving families (Orbicularia), and nymphs from six of these species, show that very small spiders resemble other small animals in having disproportionately larger central nervous systems (CNSs) relative to body mass when compared with large-bodied forms. Small spiderlings and minute adult spiders have similar relative CNS volumes. The relatively large CNS of a very small spider occupies up to 78% of the cephalothorax volume. The CNSs of very small spiders extend into their coxae, occupying as much as 26% of the profile area of the coxae of an Anapisona simoni spiderling (body mass < 0.005 mg). Such modifications occur both in species with minute adults, and in tiny spiderlings of species with large-bodied adults. In at least one such species, Leucauge mariana, the CNS of the spiderling extends into a prominent ventral bulge of the sternum. Tiny spiders also have reduced neuronal cell body diameters. The adults of nearly all orbicularian spiders weave prey capture webs, as do the spiderlings, beginning with second instar nymphs. Comparable allometric relations occur in adults of both orb-weaving and cleptoparasitic species, indicating that this behavioral difference is not reflected in differences in gross CNS allometry.     

Maternal Care and Infanticide by Males in Helvibis longicauda (Araneae: Theridiidae)

Maternal care in spiders varies from just the construction of a protective silken structure for the eggs and the selection of a safe site to place them, to a long period of association between the mother and spiderlings. Such extended care may involve the active protection from predators and parasitoids, food regurgitation, the production of trophic eggs and even matriphagy. In this study, we describe extended maternal care in Helvibis longicauda (Theridiidae) and evaluate the effectiveness of maternal protection against predators of eggs and spiderlings. We conducted experiments comparing the frequency of egg sac destruction and mortality of spiderlings in the presence and absence of mothers. We also observed the behaviour of the mother and spiderlings during prey capture events and interactions with possible predators. Helvibis longicauda females guard their egg sacs until the emergence of the young and guard the spiderlings for several instar stages, fighting possible predators, including conspecifics. We found that aggressive behaviour by females increased the survival of both eggs and spiderlings in our experiments. Intruder males were the main source of mortality in the absence of females. The benefits of maternal care for the young also include the acquisition of prey items that are captured, immobilized and pre‐digested by the mother. Effective maternal protection and the extended period of supplying food to juveniles probably contribute to the late dispersal of offspring in H. longicauda.     

Costs and Benefits of Maternal Care in a Subsocial Spider,Coelotes terrestris

The maternal social spider Coelotes terrestris demonstrates extended care towards its progeny: the mother guards its egg sac for 3–4 weeks, then stays with its young from the time of their emergence until their dispersal about 1 month later. The present investigation evaluates the adaptiveness of these maternal behaviours by comparing the fitness of females performing them with that of females separated from the egg sac or the spiderlings. By protecting their egg sacs from predation and parasites, and by pursuing this task while supplying the young with food, mothers enhance the survival rate and the development of many of their spiderlings. The costs linked with these activities, estimated by the ability to produce another clutch, appear variable according to the stage in the reproductive cycle. In such terms, the egg sac guarding appears to have a low cost in relation to the care given to the spiderlings.     

Seasonal changes in resource allocation within an individual offspring of the wolf spider, Pardosa pseudoannulata (Araneae: Lycosidae)

Abstract.  Seasonal changes in resource investment into individual offspring are well documented, but no attention has been paid to the allocation of the invested resource among the body parts of the offspring. In the present study, seasonal changes in the absolute and relative sizes in a spiderling of the wolf spider Pardosa pseudoannulata Boes. et Str. are investigated, and the relationship between spiderling body size and the ability to moult during a food shortage is clarified. Spiderlings that emerge in November have a significantly larger cephalothorax and abdomen than those that emerge in June. In addition, the abdomen–cephalothorax size ratio is significantly greater in November than in June offspring. Under limited food availability conditions, only 40% of spiderlings moult. Nymphs that do moult have a significantly larger cephalothorax, abdomen and abdomen–cephalothorax size ratio than nymphs that do not moult. Thus, both the quantity of resources invested in the cephalothorax and abdomen of a spiderling and the proportional allocation of resources between the two body parts change seasonally in Pa. pseudoannulata ; alteration of the resource allocation occurs in late autumn. Larger spiderlings of Pa. pseudoannulata that emerge in late autumn would be able to develop into advanced instars even during food shortage, and therefore may have better overwintering survival rates.     

寡居种悦目金蛛若蛛群居机制初步研究

对寡居种悦目金蛛Argiope amoena若蛛群居生活对其结网的影响、温度和种群大小对若蛛存活率的影响、若蛛对限制性空间的利用及其扩散方式进行了观察和研究.结果表明,若蛛从卵袋出蛰后不经历群居扩散即具备结网能力,可结完整网,也具备在扩散前不进食水存活的能力;在变温条件下若蛛存活率要远远高于室温条件下的存活率,而在室温条件下若蛛开始死亡和半数死亡的时间比其聚居期长得多;若蛛主要通过群体空中扩散的方式进行扩散,可以减小能耗与敌害的威胁;不同种群大小的若蛛在限制性空间的分布均从第5d左右开始发生较大变化的,这与在空间不受限的条件下若蛛在第5d开始扩散的结果是一致的.据此我们推断:个体扩散代价最小化(包括御敌和能耗)和维持种群较高的生存率可能是寡居种悦目金蛛若蛛群居生活的主要原因.     

Female fecundity and offspring survival are not increased through sexual cannibalism in the spider Larinioides sclopetarius

Many hypotheses explaining the evolution and maintenance of sexual cannibalism incorporate the nutritional aspect of the consumption of males. Most studies have focused on a fecundity advantage through consumption of a male; however, recent studies have raised the intriguing possibility that consumption of a male may also affect offspring quality. In particular, recent studies suggest prolonged survival for offspring from sexually cannibalistic females. Here, we measured the protein and lipid content of males compared to insect prey (crickets), quantified female nutrient intake of both prey types and finally assessed how sexual cannibalism affects female fecundity and spiderling quality in the orb‐web spider Larinioides sclopetarius. We found no evidence that sexual cannibalism increased fecundity when compared to a female control group fed a cricket. Contrary to previous studies, spiderlings from females fed a male showed reduced survival under food deprivation compared to spiderlings from the control group. Offspring from females fed a male also tended to begin web construction sooner. The low lipid content of males compared to crickets may have reduced offspring survival duration. Whether additional proteins obtained through consumption of a male translate to enhanced silk production in offspring requires further investigation.     

Kinship Influences Cannibalism in the Wolf Spider, Pardosa milvina

Recent studies have called into question the role of Wright's coefficient of relatedness (r) in the interactions among relatives. Kin selection theory predicts a positive relationship between relatedness and frequency of altruistic acts, but a number of researchers have reported the opposite relationship. I used a lycosid spider (Pardosa milvina) to test the hypothesis that genetic relatedness would affect the propensity of a cannibalistic species to prey on genetic relatives. I considered lack of predation to be a form of altruism where the predator incurs a cost (loss of a meal) that benefits potential prey. Specifically, I questioned whether direct genetic offspring would be avoided as prey items and whether the sex or reproductive condition of a cannibalistic predator would affect the likelihood of predation on conspecific juveniles. As predicted by kin selection theory, spiderling mothers ate significantly fewer of their own offspring than they did of nonkin spiderlings of the same age. Adult virgin female and adult male spiders ate significantly more spiders than females that had recently carried spiderlings. Females with egg sacs consumed significantly fewer spiderlings than did virgin female spiders. These findings support Hamilton's rule and suggest that, in some systems, genetic relatedness plays a strong role in governing altruistic behavior toward relatives.     

Infanticide and Maternal Offspring Defence in European Rabbits under Natural Breeding Conditions

Infanticide is an important source of mortality of dependent offspring in several mammal species, whereas female conspecifics are often the perpetrators. This has led to maternal counter‐strategies, such as the defence of the nests. However, cases of infanticide are hard to detect in the field, and studies on maternal offspring defence behaviour under natural breeding conditions are scarce. We conducted such a study on the European rabbit (Oryctolagus cuniculus), which is usually considered to show low maternal care. The study was carried out over 5 yr on a field enclosure population. We (1) studied infanticide rates and the impact of potential determinants: the group density and age structure of the females’ rank hierarchy within the groups; we used the latter as an estimator of social group stability. We (2) studied if mothers defend their breeding burrow against approaching, potentially infanticidal females. Overall, we recorded infanticide in 5% of all litters; infanticide was the cause in 12% of cases of litter mortality. The proportion of infanticide was 7% higher in groups where same‐age females occupied successive rank positions than in groups where the females’ rank hierarchy had a more heterogeneous and linear age structure. We hypothesize that social instability in the former groups was the reason for the increased infanticide risk. Infanticide rates were not correlated with group density and did not differ among mothers with different social ranks. Infanticide occurred exclusively during the first 10 d after parturition. During this time, mothers stayed closer to their breeding burrows than shortly before parturition or during later lactation. Moreover, mothers were more aggressive against other females in proximity to their breeding burrow than in more distant areas. We suggest that the pattern of spacing behaviour and intrasexual aggression of rabbit mothers are an adaptive response to the occurrence of female infanticidal behaviour.     

Physiological costs during the first maternal care in the wolf spider Pardosa saltans (Araneae,Lycosidae)

Many arachnids like other terrestrial arthropods, provide extensive maternal care. Few studies have quantified the underlying physiological costs of maternal care. We investigated how maternal care affects the free-moving wolf spider’s (Pardosa saltans) energy requirements. We described in detail their basic reproduction biology (i.e. carrying cocoon and young) and we evaluated the variation in the females’ energy reserves during maternal care. Our results show that mothers guard eggs until hatching and then guard their spiderlings for 27–30 more days. Laboratory observations indicated that spiderlings start leaving the maternal abdomen gradually 5–7 days after hatching. Females carry an egg sac (cocoon) that can weigh up to 77% of their post-reproduction weight and carry young that weigh 87–100% of their body mass. Females lost weight over time despite regular food intake, while carrying cocoon and young; but their weights increased gradually during the dispersal of young. The contributions of proteins, glucose and triglycerides to maintain females’ energy were calculated. Their energetic state varied during maternal care, in particular lipid levels declined, during the care of spiderlings when the females’ predatory behaviour was inhibited. Our results show that the maternal care provided by P. saltans females is particularly costly physiologically, during the 30 days following egg sac formation and development of spiderlings, even when food is available.     

Drop and swing dispersal behavior of a tropical wandering spider: experiments and numerical model

Summary When exposed to certain air flows spiderlings of the wandering spider Cupiennius getazi (Ctenidae) drop from their dwelling plant and swing in the wind from a gradually lengthening dragline. If body contact is made with a nearby substrate the spiderling detaches. We refer to this form of aerial dispersal as the drop and swing dispersal behavior (DASDB). The dragline being only up to about 70 cm long and only rarely ruptured by the drag forces of the wind, this is a close range type of dispersal as opposed to the ballooning known for many other species of spiders. DASDB is readily elicited in spiderlings at an age of ca. 9 days (outside egg sac). At this age their mass is 1.26 ±0.35 mg and their yolk usually depleted. They then start to catch prey and escape from the unfavorable conditions in the small space around the egg sac where hundreds of spiderlings compete. Air flow rates effectively eliciting DASDB in the laboratory are between 0.2 m/s and 1.5 m/s. The number of spiderlings showing DASDB increases considerably if the air flow is turbulent as opposed to laminar. A numerical model defining the window within which DASDB is supported mechanically was developed from theoretical considerations. Taking the effective wind speeds and the mechanical properties of the dragline, the model accounts very well for the fact that actual rupture of the dragline was observed only rarely in C. Getazi. Other features of the DASDB are also correctly predicted. The model is not only applicable to DASDB but also to the drop and swing preballooning behavior known to occur in several other species of spiders.Abbreviations DASDB drop and swing dispersal behavior     

Relationship between reproductive resource allocation and resource capacity in the matriphagous spider, <Emphasis Type="Italic">Chiracanthium japonicum </Emphasis>(Araneae: Clubionidae)

Females of the Japanese foliage spider, Chiracanthium japonicum, are eaten by their offspring at the end of the maternal care period. To examine the patterns of allocation of maternal investment to their offspring associated with female resource capacity, the amounts of female body reserves accumulated before oviposition, reproductive components at the egg-production phase and those at the matriphagy phase were measured using an artificial breeding nest. Regardless of size, female bodies were completely consumed by the offspring, and larger females, i.e. those having larger reserves, produced a larger number of offspring, but not larger offspring. Furthermore, the proportion of reserves allocated to egg production was not affected by the total amount of the reserves, which indicated that the females systematically divided the resources for individual offspring between egg yolk and food for the growth and survival of the offspring. These results suggest that C. japonicum females adjust egg production to their own resource capacity so that they can achieve an investment per individual offspring which is not dependent on resource capacity. Electronic Publication     

Variation in life-history characteristics over a clinal gradient in three populations of a communal orb-weaving spider

Summary Adult size and fecundity, total reproductive biomass, egg size, and spiderling size and resistance to desiccation were compared in three populations of the communal orb-weaving spider,Metepeira spinipes, in Mexico. In a desert population and in one from a moist tropical forest adult size, fecundity, and total reproductive biomass were similar, but were markedly smaller than in a climatologically intermediate agricultural habitat. Egg size and protein content were greater in the desert and agricultural habitats than in the moist forest, but spiderling size increased from desert to agricultural to moist forest populations. Desert spiderlings survived significantly longer than moist forest spiderlings at all humidity treatments over a 10% to 100% range. An explanation for these results is proposed based on apparent differences in energy allocation and expenditure which arise from the distinct climate, colony structure, and prey and space availability in the three habitats.     

Major benefits of guarding behavior in subsocial bees: implications for social evolution

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.     

Social behaviour of spitting spiders (Araneae: Scytodidae) from Singapore

The social organization of Scytodes sp., a spitting spider from Singapore, was investigated in the field and in the laboratory. The characteristics considered were colony structure, maternal care, prey capture, prey sharing, adults provisioning with prey and how feeding regime influences the level of tolerance expressed by juveniles that were living together. The characteristics of this species were compared with those of other Scytodes species that have been studied previously. Based on our findings, we conclude that in this species, social organization is 'periodic-social', with extended maternal care, this being a type of 'subsociality'. The females of Scytodes sp. wrapped their eggs in a silk egg sac and used their chelicerae to hold the eggs until they hatched. Spiderlings tended to remain in their mother's web until the fourth instar (i.e. after the third moult) but they were also highly aggressive and cannibalistic towards conspecifics after the third instar. Females carried prey to their spiderlings and either fed alongside the brood or left them to feed alone. Spiderlings also captured large prey cooperatively by spitting onto, biting and feeding on it together. The occurrence of cannibalism among her juveniles did not appear to be influenced by whether the adult female was present or not, nor did it appear to be influenced by prey abundance.     

Variation in group cohesion during the maternal social period of a subsocial spider

During the maternal social period, Amaurobius ferox spiderlings (Araneae: Amaurobiidae) show mutual tolerance, group cohesion and cooperation in prey capturing, which are recognized as the main characteristics in the evolution of spider sociality. Measuring spatial volume occupied by the spiderlings within the maternal web, this study investigated variation in group cohesion over the maternal social period, from emergence to dispersal. The results showed that the spatial volume of spiderlings varied greatly during the maternal social period and was associated with the development of the spiderlings. Strong group cohesion appeared to be related to maternal food provision, trophic egg laying and matriphagy. An increase of the spatial volume was obviously observed after matriphagy. The experiment revealed that group cohesion decreased after the second molt. A compact group of spiderlings should facilitate maternal food production at the prompted time, and reduced group cohesion after the second molt might lead to eventual dispersal of A. ferox spiderlings.     

COOPERATIVE BREEDING FAVORS MATERNAL INVESTMENT IN SIZE OVER NUMBER OF EGGS IN SPIDERS

The transition to cooperative breeding may alter maternal investment strategies depending on density of breeders, extent of reproductive skew, and allo‐maternal care. Change in optimal investment from solitary to cooperative breeding can be investigated by comparing social species with nonsocial congeners. We tested two hypotheses in a mainly semelparous system: that social, cooperative breeders, compared to subsocial, solitarily breeding congeners, (1) lay fewer and larger eggs because larger offspring compete better for limited resources and become reproducers; (2) induce egg size variation within clutches as a bet‐hedging strategy to ensure that some offspring become reproducers. Within two spider genera, Anelosimus and Stegodyphus, we compared species from similar habitats and augmented the results with a mini‐meta‐analysis of egg numbers depicted in phylogenies. We found that social species indeed laid fewer, larger eggs than subsocials, while egg size variation was low overall, giving no support for bet‐hedging. We propose that the transition to cooperative breeding selects for producing few, large offspring because reproductive skew and high density of breeders and young create competition for resources and reproduction. Convergent evolution has shaped maternal strategies similarly in phylogenetically distant species and directed cooperatively breeding spiders to invest in quality rather than quantity of offspring.     

Paternal levels of DNA damage in spermatozoa and maternal parity influence offspring mortality in an endangered ungulate

Understanding which factors influence offspring mortality rates is a major challenge since it influences population dynamics and may constrain the chances of recovery among endangered species. Most studies have focused on the effects of maternal and environmental factors, but little is known about paternal factors. Among most polygynous mammals, males only contribute the haploid genome to their offspring, but the possibility that sperm DNA integrity may influence offspring survival has not been explored. We examined several maternal, paternal and individual factors that may influence offspring survival in an endangered species (Gazella cuvieri). Levels of sperm DNA damage had the largest impact upon offspring mortality rates, followed by maternal parity. In addition, there was a significant interaction between these two variables, so that offspring born to primiparous mothers were more likely to die if their father had high levels of sperm DNA damage, but this was not the case among multiparous mothers. Thus, multiparous mothers seem to protect their offspring from the deleterious effects of sperm DNA damage. Since levels of sperm DNA damage seem to be higher among endangered species, more attention should be paid to the impact of this largely ignored factor among the viability of endangered species.